Mollusca: Olividae) from the Southwestern Atlantic
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Zootaxa 2889: 1–34 (2011) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2011 · Magnolia Press ISSN 1175-5334 (online edition) A revision of the genus Olivancillaria (Mollusca: Olividae) from the southwestern Atlantic VAL ER IA TE SO 1 & GUIDO PASTORINO Museo Argentino de Ciencias Naturales Av. Angel Gallardo 470, Lab. 57 C1405DJR Buenos Aires, Argentina 1Corresponding author. E-mail: [email protected] Table of contents Abstract . 1 Introduction . 2 Material and methods . 2 Results . 4 Morphometric analysis . 4 Systematics . 6 Class Gastropoda Cuvier, 1798 . 6 Subclass Orthogastropoda Ponder & Lindberg, 1997 . 6 Superorder Caenogastropoda Cox, 1959 . 6 Order Sorbeoconcha Ponder & Lindberg, 1997 . 6 Infraorder Neogastropoda Thiele, 1929. 6 Superfamiliy Muricoidea Rafinesque, 1815 . 6 Family Olividae Latreille, 1825 . 6 Subfamily Olivinae Latreille, 1825 . 6 Genus Olivancillaria d´Orbigny, 1840 . 7 Olivancillaria urceus (Röding, 1798) . 7 Olivancillaria vesica (Gmelin, 1791) . 10 Olivancillaria auricularia (Lamarck, 1811) . 13 Olivancillaria contortuplicata (Reeve, 1850) . 15 Olivancillaria deshayesiana (Ducros de Saint Germain, 1857) . 16 Olivancillaria orbignyi (Marrat, 1868) . 20 Olivancillaria teaguei Klappenbach, 1964 . 24 Olivancillaria carcellesi Klappenbach, 1965 . 24 Species excluded from Olivancillaria . 28 Discussion . 29 Key of the species of Olivancillaria . 30 Acknowledgements . 30 References . 31 Abstract The genus Olivancillaria is revised and restricted to eight living species: O. deshayesiana, O. carcellesi, O. urceus, O. contortuplicata, O. orbignyi, O. teaguei, O. auricularia and O. vesica. The last two species were previously considered as subspecies. The name O. uretai is a junior synonym of O. orbignyi. The genus Olivancillaria is endemic to the south- western Atlantic, ranging from Bahia state, Brazil (~12°S) to Chubut province (42°37’S), Argentina, intertidal to 70 m depth. It is a clear representative of the Argentine Malacological province. Redescriptions and re-illustration of types were done for each species. Shell ultrastructure, radulae, penes, siphon papillae and egg capsules are described and illustrated by SEM images. A geometric morphometric analysis was carried out to confirm shell variation on size and shape among species. The geographic distribution of each species is provided based on field observations as well as on museum records. Key words: Argentina, Gastropoda, geometric morphometric analysis, Neogastropoda, Olivancillaria, Olividae Accepted by M. deMaintenon: 11 Apr. 2011; published: 24 May 2011 1 Introduction The family Olividae Latreille, 1825 includes carnivorous or detritivorous semi-infaunal marine gastropods of mod- erate size (Smith 1998). They inhabit nearshore waters in temperate and tropical regions. Thirty species of Olividae are reported from South America, from the genera Olivancillaria d´Orbigny, 1840, Oliva Bruguière, 1789, Olivella Swainson, 1831, Ancilla Lamarck, 1799, Jaspidella Olsson, 1956, Agaronia Gray, 1839 and Amalda Adams & Adams, 1853 (see e.g. Absalão 2000, Absalão & Pimenta 2003, Pastorino 2003, Rios 2009). Among the regional taxonomic treatments specifically of the genus Olivancillaria, Marcus & Marcus (1959) described the anatomy of O. urceus (as O. brasiliensis) and O. auricularia (under the genus Lintricula). Later Klappenbach (1964, 1965, 1966) published a series of papers where he described three species and discussed the status of two others belonging to this genus. The last account of Olivancillaria species was done by Thomé (1966), who described O. buckuporum from southern Brazil, considered later as a local variation of O. carcellesi (Rios 1970, Thoméet al. 2004). In a very interesting approach, Tursch (1988) studied several protoconch parameters of some species of Olivi- dae and confirmed supraspecific differences among several genera including Olivancillaria. Carranza & Norbis (2005) have documented the variation among the shells of O. urceus from several Southwestern Atlantic localities. Recently, Teso et al. (2010) studied the phenotypic variation of shell size and shape of O. carcellesi from four Southwestern localities using geometric morphometric methods. The aim of this paper is to revise and redescribe the living species of Olivancillaria including examination of type material, corroborated ranges, together with SEM illustrations of shell ultrastructure, radulae, penes, siphonal papillae and, when available, egg capsules. In addition, a geometric morphometric analysis was carried out to con- firm shell variation in size and shape among species. Material and methods Live specimens were collected from several localities along the Argentine coast: intertidally from San Clemente del Tuyú (36°22′S, 56°42′W) and Santa Teresita (36°32′S, 56°40′W); by bottom trawling in 4–12 m depth from Villa Gesell (37°15′S, 56°58′W), Mar Chiquita (37°44′S, 57°24′W) and Mar del Plata (38°02′S, 57°31′W); by SCUBA diving in 6–20 m depth from Puerto Lobos (42°00′S, 65°04′W) and Punta Pardelas (42°37'S, 64°15'O) (Figure 1). Other specimens analyzed are housed at the following institutions: Museu Nacional de Rio de Janeiro (MNRJ), Museu de Zoologia da Universidade de São Paulo (MZUSP) and Museu Oceanográfico “Profesor Eliézer de Carvalho Rios”, Rio Grande (MORG), Brazil; Museo Nacional de Historia Natural (MNHNM) and Barattini’s collection at the Museo Zoológico Municipal “Dámaso Larrañaga”, both in Montevideo, Uruguay; Museo Argen- tino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires (MACN-In) and Museo de La Plata (MLP), Argentina; The Natural History Museum, London (NHMUK) and National Museums Liverpool (NML); Muséum d´Histoire naturelle, Genève (MHNG); Natuurhistorisch Museum, Rotterdam (NMR), The Netherlands; Muséum national d´Histoire naturelle, Paris (MNHN), the Zoological Institution, St-Petersburg, Russia, Museum of Natural History Vienna (NHMW), Austria and Universidade de Santiago de Compostela (USC), Spain. All shells were photographed using a Nikon D100 camera with a Micro Nikkor lens and digitally processed with the appropriate software. All known citations of living records for the genus Olivancillaria in the literature are included on the syn- onymic lists (See Table 2 for index). Dissections were performed on ethanol-preserved specimens for study of the gross anatomy of the anterior ali- mentary system, and the pallial portions of the male and female reproductive systems. Radulae were cleaned with NaClO and sonicated in an ultrasonic cleaner from about five males and five females of each species. Shell ultra- structure was studied using freshly fractured collabral sections taken from the central portion of the lip on the last whorl. The shell thickness was divided into four groups: thickest (~2.3 mm), thick (~1 mm), medium (~0.6 mm) and thinnest (~0.4 mm). SEM photographs were taken using a Philips XL30 at MACN. Geometric morphometric analysis was performed on twenty specimens of each species (Table 1). Twelve land- marks representative of the shell (sensu Rohlf 1996, Roopnarine & Vermeij 2000) were chosen on apertural view 2 · Zootaxa 2889 © 2011 Magnolia Press TESO & PASTORINO (Figure 2); for descriptions of each landmark see Teso et al. (2010). The landmark data were used to calculate the specimen size and to describe the shape variation among species. Size was calculated as the square root of the summed squared distances of a specimen’s landmarks to their geometric centroid: CS = [Σ (di) 2]1/2, where CS is centroid size and di is the distance of landmarks to their centroid. The description of shape was carried out by rela- tive warp analysis (RWA) (Bookstein 1991) performed with the tpsRelw (Rohlf 2005). This method is comparable to principal component analysis, which summarizes a specimen shape as a deformation from a common reference. The relative warps were computed with the scaling option α = 0 which weights all landmarks equally, for studying differences in shell shape among samples (Rohlf et al. 1996). In addition, the interspecific variation among groups was tested with a multivariate analysis of variance (MANOVA) of partial warp scores (Zelditch et al. 2004). The posterior categorization was revealed by canonical variates analysis (CVA). We also used a discriminant function analysis (DFA) with the RWs (Relative Warps) and CS (centroid size) to test differences between groups. Allomet- ric variation was tested with a multiple regression model with the relationships between RWs and CS and with a covariance analysis (ANCOVA). Differences in centroid size distributions among groups were tested with one-way ANOVA and post hoc Tukey HSD test (Sokal & Rohlf 1995). Normality was tested with Lilliefors test. Statistical analysis was carried out with Statistic v. 6.0. When homogeneity of variances could not be achieved by transforma- tion, data were analyzed nonetheless, since analysis of variance is robust for departure from this assumption when sizes of samples are equal (Underwood 1997). Results were, however, interpreted with caution by judging signifi- cance more conservatively (α = 0.01). FIGURE 1. Map showing the sampled localities. OLIVANCILLARIA FROM SOUTHWESTERN ATLANTIC Zootaxa 2889 © 2011 Magnolia Press · 3 FIGURE 2. Apertural view of Olivancillaria carcellesi (MACN-In 37505) showing the homologous landmarks used in the geometric morphometric analysis. Scale bar = 1 cm. Results Morphometric analysis The geometric morphometric method by means of MANOVA and posterior CVA showed significant differences among groups (Wilks’ λ = 0.00001, χ2 = 1464.4501, P < 2.22045 exp-16). The study revealed eight morphotypes 4 · Zootaxa 2889 © 2011 Magnolia Press TESO & PASTORINO (Figure 3). We found that 100% of the original cases were cross-validated correctly into their species groups based on the Mahalanobis distance in the space defined by.